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Introduction


The decrease in stratospheric ozone has prompted renewed efforts in assessing the potential damage to plant and animal life due to enhanced levels of solar Ultraviolet-B (UV-B, 280-320 nm) radiation (Caldwell 1971, 1998; Madronich et al., 1998). The effect of UV-B enhancements on plants includes reduction in yield and quality, alteration in species competition, decrease in photosynthetic activity, susceptibility to disease, and changes in plant structure and pigmentation (Tevini and Teramura 1989; Bornman 1989; Teramura and Sullivan 1991). Some species show sensitivity to present levels of UV-B radiation while others are apparently unaffected by rather massive UV enhancements (Becwar et al., 1982). This issue is complicated further by reports of equally large response differences among cultivars of a species (Biggs et al., 1981; Teramura and Murali 1986). About two-thirds of some 300 species and cultivars tested appear to be susceptible to damage from increased UV-B radiation. Crops such as soybean, winter wheat, cotton, and corn are susceptible to damage from increased UV-B radiation. All effects of elevated UV-B on plants should be considered in the context of other factors such as water stress, increased atmospheric CO2, tropospheric air pollution, and temperature. The effects of UV-B on plants have been studied mostly under growth chamber, greenhouse, while a few experiments conducted under field conditions (Krupa, 1989). There are also few studies that have examined the joint effects of UV-B and other stress factors on plant response. The effect of UV-B on plant growth and productivity varies seasonally and is affected by microclimate and soil fertility. For instance, soybeans are less susceptible to UV-B radiation under water stress or mineral deficiency, but sensitivity increases under low levels of visible radiation (Teramura, 1983). Continued studies over many growing seasons are crucial in any UV-B impact assessment of agricultural productivity. 

Methodology for the assessment of UV-B effects on plants

In this section only a very brief discussion of methods available for studying the effects of UV-B on plants (summarized from Krupa, 1989) is provided. The measurements and physical simulation of UV-B radiation in the growth chamber, greenhouse or under ambient field conditions is not straightforward. Table 1 gives a summery of methods used for the examining the effects of UV-B on plants. The general principle in the experiments to determine the effects of UV-B on plants involves the use of a UV source (a lamp) coupled with different types of filters to exclude bands of UV wavelength not desired in the experiment. The intensity of UV is varied by changing the height between the lamp source and the plant canopy. Because different biological processes exhibit different degrees of sensitivity to different wavelengths of UV-B, a mathematical response function, the action spectrum, must be used as a weighting factor to adjust the measured UV-B flux. The various sources of uncertainties in calculating biologically effective UV-B flux should be considered.

Table 1

Summery of methods used to determine the effects of UV-B on plants


Methods

Reference


Greenhouse:
UV lamps and selective wavelength filters, Westinghouse FS-40 sun lamp frames with cellulose acetate or Mylar type S filters

Dumpert & Knacker (1985)
Mirecki & Teramura (1984)
Growth chamber:
UV-B lamps, simulated PAR (photosynthetic active radiation) and selective wavelength cut-off filters

Tevini & Iwanzik (1986)
Field exposure:
FS-40 sun lamps coupled with Aclar, Mylar and cellulose acetate filters, Modulated fluorescent lamp system for  supplementing natural UV-B

Becwar et al. (1982)
Lydon et al. (1986)
Caldwell et al. (1983a)

UV-B effects on plants

Agricultural scientists have responded with a series of pioneering investigations on the effect of artificial and solar UV radiation upon plant growth and development. A great variety of physiological and morphological plant responses to UV radiation have been subsequently demonstrated over the past years. Most of these experiments, however, have employed UV lamps which usually emit radiation quite unlike the radiation present in the normal terrestrial solar spectrum. The importance of solar angle, atmospheric turbidity, elevation above the sea level, cloud cover, total atmospheric ozone column, and the UV albedo of the earth's surface with respect to the total UV irradiation intensity and wavelength composition should be considered in UV radiation of natural environments. Though not all the plant responses demonstrated as the result of UV radiation are considered as damaging or disadvantageous for the plant; the majority of evidence indicates that UV irradiation is usually detrimental, particularly UV-B irradiation (Caldwell, 1971). In this section a summery of the UV-B effects on crops from the literature will be presented. The growth of many plant species is reduced by enhanced levels of UV-B radiation (Teramura et al., 1989). The enhanced UV-B radiation generally has negative impacts on growth, yield and quality of some crop plants such as soybean, winter wheat, rice, sorghum, cotton and corn. The response varies with different plant species. Some are very sensitive and some are least sensitive. With enhanced UV-B radiation photosynthesis decreases, plant height and leaf area decrease, dry matter production, yield and quality reduces in many crops. In the study conducted by Tevini et al. (1991b) plant height, leaf area, and the dry weight of sunflower, corn, and rye seedlings were significantly reduced with enhanced UV-B radiation. Rice is among the most important crop plants in the world. Sixteen rice cultivars from several different geographical regions were grown in greenhouses with supplemental levels of UV-B radiation (Teramura et al., 1991). Alterations in biomass, morphology, and maximum photosynthesis were determined. Approximately one-third of all cultivars tested showed a statistically significant decrease in total biomass with increased UV-B radiation. For these sensitive cultivars, leaf area and tiller number were also significantly reduced. Photosynthetic capacity, as determined by oxygen evolution, declined for some cultivars. In a six year field study of a UV-sensitive soybean, Teramura et al. (1990) presented a statistically significant 19%-25% reduction in seed yield in five of the six years under a 25% ozone reduction level. 

Reference

Becwar, M.R., F.D. Morre III, and M.J. Bureke. 1982. Effects of depletion and enhancement of ultraviolet-B (280-315nm) radiation on plants grown at 3000 m elevation. J. Amer. Soc. Hort. Sci. 107:771-779.

Biggs, R.H., S.V. Kossuth, and A.H. Teramura. 1981. Response of 19 cultivars of soybeans to ultraviolet-B irradiance. Physiol. Plant. 53:19-26.

Bornman, J.F. 1989. Target sites of UV-B radiation in photosynthesis of higher plants. J. Photochem. Photobiol. B: Biol. 4:145-158.

Caldwell, M.M. 1971. Solar UV irradiation and growth and development of higher plants. p. 131-177. In A.C. Giese (ed.) Photophysiology, Volume 4.

Caldwell, M.M., L.O. Bjorn, J.F. Bornman, S.D. Flint, G. Kulandaivelu, A.H. Teramura, and M. Tevini. 1998. 

Effects of increased solar ultraviolet radiation on terestrial ecosystems. J. Photochem. Photobiol. B: Biol. 46(1-3):40-52.

Krupa, S.V., R.N. Kickert. 1989. The greenhouse effect impacts of ultraviolet-B (UV- B) radiation, carbon dioxide (CO), and ozone (O) on vegetation. Environmental Pollution 61:263-393.

Madronich, S., R.L. McKenzie, L.O. Bjorn, and M.M. Caldwell. 1998. Changes in biologically active ultraviolet radiation reaching the Earth's surface. J. Photochem. Photobiol. B: Biol. 46(1-3):5-19.

Teramura, A.H., M. Tevini and W. Iwanzik. 1983. Effects of ultraviolet-B irradiance on plants during mild water stress. I. Effects on diurnal stomatal resistance. Physiol Plant. 57:175-180.

Teramura, A.H., and N.S. Murali. 1986. Intraspecific differences in growth and yield of soybean exposed to ultraviolet-B radiation under greenhouse and field conditions. Environ. and Experi. Botany. 26:89-95.

Teramura, A.H., J.H.Sullivan, and J.Lydon. 1990. Effects of solar UV-B radiation on Soybean yield and seed quality: a six-year field study. Physiologia Plantarum.80: 5-11.

Teramura, A.H., and J.H. Sullivan. 1991. Potential effects of increased solar UV-B on global plant productivity. p. 625-634. In E. Riklis (ed.) Photobiology, Plenum Press, New York.

Tevini. M., and A.H. Teramura. 1989. UV-B effects on terrestrial plants. Photochem. Photobiol. 50:479-487.

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